Development of telotrophic‐meroistic ovarioles of polyphage beetles with special reference to the formation of nutritive cords

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Abstract
Oocytes and nurse tissue of Bruchidius differentiate from germ cells during the extended period of pupal development (7.2 ± 0.6 days). A system of 15 pupal stages correlates ovarian development with changes in pigmentation of the eyes, maxillae, alae and tarsalia. The ovarioles grow in length at a constant rate, though their width does not change. A differentiating zone, consisting of germ cells and the basal layer of interstitial cells, arises at the base of the tropharium and separates presumptive oocytes and nurse cells. Early in pupal development the germ cells are arranged in primary syncytia with the cells connected by persisting intercellular bridges filled with fusomal material, never with larger particles, such as mitochondria. At later stages membrane disintegration changes the primary syncytium into a secondary one including all nurse cell nuclei. Nutritive cords are first noticeable when differentiation of oocytes and nurse cells starts. The cords seem to be of primary origin, i.e., they are connections between sister cells which become elongated as these cells are separated during growth. This is indicated by the persistence of intercellular bridges which are sometimes found as part of the membrane of growing nutritive cords connecting young oocytes with the nurse cell syncytium.